Zootaxa: The correct use of the name Scytalopus speluncae ....

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1271 Accepted by S. Olson: 22 Jun 2006; published: 24 Jul. 2006 37 ZOOTAXA ISSN 1175-5326 (print edition) ISSN 1175-5334 (online edition) Copyright © 2006 Magnolia Press Zootaxa 1271: 3756 (2006) www.mapress.com/zootaxa/ The correct use of the name Scytalopus speluncae (Ménétriés, 1835), and the description of a new species of Brazilian tapaculo (Aves: Passeriformes: Rhinocryptidae) MARCOS A. RAPOSO 1 , RENATA STOPIGLIA 1 , VLADIMIR LOSKOT 2 & GUY M. KIRWAN 3 1 Museu Nacional, Departamento de Vertebrados, Setor de Ornitologia, Quinta da Boa Vista, 20940-040, Rio de Janeiro, RJ, Brazil. E-mail: [email protected] 2 Department of Ornithology, Zoological Institute, Russian Academy of Science,. St. Petersburg, 199034, Russia 3 74 Waddington Street, Norwich NR2 4JS, UK. Abstract Scytalopus speluncae was described from a specimen taken at São João del Rei, Minas Gerais, Brazil. Ménétriés described it as having the throat and the central breast evidently whitish, but since then this name has been rather surprisingly attributed to the uniformly slate gray Mouse-colored Tapaculo of the eastern coastal Atlantic Forest of Brazil. Contrary to previous findings, our analysis of the holotype, two new specimens from the type locality (topotypes) and original description of Scytalopus speluncae indicates that this name must be applied to the pale gray form with rufous- barred thighs that is found in Minas Gerais. This species is apparently very common in São João del Rei. This paper redescribes Scytalopus speluncae, correcting this name application, and describing, as a new species, populations from the Serra do Mar and vicinity, previously misidentified as S. speluncae. Key words: Scytalopus speluncae, new species, Brazil, taxonomy Resumo Scytalopus speluncae foi descrita com base em um espécime proveniente de São João del Rei, Minas Gerais, Brasil. Ménétriés descreveu essa espécie como tendo a garganta e o centro do peito “evidentemente esbranquiçados” mas, desde então, esse nome tem sido surpreendentemente atribuído ao “tapaculo-preto” da floresta atlântica da costa brasileira. Contrariamente aos estudos anteriores, nossa análise do holótipo, de dois espécimes recentemente coletados na localidade tipo (topótipos) e da descrição original de Scytalopus speluncae indicam que esse nome deve ser atribuído à forma cinza clara com “calções” ocres barrados que é encontrada em Minas Gerais, a

Transcript of Zootaxa: The correct use of the name Scytalopus speluncae ....

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1271

Accepted by S. Olson: 22 Jun 2006; published: 24 Jul. 2006 37

ZOOTAXAISSN 1175-5326 (print edition)

ISSN 1175-5334 (online edition)Copyright © 2006 Magnolia Press

Zootaxa 1271: 37–56 (2006) www.mapress.com/zootaxa/

The correct use of the name Scytalopus speluncae (Ménétriés, 1835), and the description of a new species of Brazilian tapaculo (Aves: Passeriformes: Rhinocryptidae)

MARCOS A. RAPOSO1, RENATA STOPIGLIA1, VLADIMIR LOSKOT2 & GUY M.

KIRWAN 3

1Museu Nacional, Departamento de Vertebrados, Setor de Ornitologia, Quinta da Boa Vista, 20940-040, Rio de Janeiro, RJ, Brazil. E-mail: [email protected]

2Department of Ornithology, Zoological Institute, Russian Academy of Science,. St. Petersburg, 199034, Russia374 Waddington Street, Norwich NR2 4JS, UK.

Abstract

Scytalopus speluncae was described from a specimen taken at São João del Rei, Minas Gerais,Brazil. Ménétriés described it as having the throat and the central breast evidently whitish, but sincethen this name has been rather surprisingly attributed to the uniformly slate gray Mouse-coloredTapaculo of the eastern coastal Atlantic Forest of Brazil. Contrary to previous findings, our analysisof the holotype, two new specimens from the type locality (topotypes) and original description ofScytalopus speluncae indicates that this name must be applied to the pale gray form with rufous-barred thighs that is found in Minas Gerais. This species is apparently very common in São João delRei. This paper redescribes Scytalopus speluncae, correcting this name application, and describing,as a new species, populations from the Serra do Mar and vicinity, previously misidentified as S.speluncae.

Key words: Scytalopus speluncae, new species, Brazil, taxonomy

Resumo

Scytalopus speluncae foi descrita com base em um espécime proveniente de São João del Rei,Minas Gerais, Brasil. Ménétriés descreveu essa espécie como tendo a garganta e o centro do peito“evidentemente esbranquiçados” mas, desde então, esse nome tem sido surpreendentementeatribuído ao “tapaculo-preto” da floresta atlântica da costa brasileira. Contrariamente aos estudosanteriores, nossa análise do holótipo, de dois espécimes recentemente coletados na localidade tipo(topótipos) e da descrição original de Scytalopus speluncae indicam que esse nome deve seratribuído à forma cinza clara com “calções” ocres barrados que é encontrada em Minas Gerais, a

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1271ZOOTAXA qual é muito comum em São João del Rei. Este artigo redescreve Scytalopus speluncae, corrigindo

a aplicação desse nome e descrevendo, como uma nova espécie, a população da Serra do Mar eadjacências, que antes era erradamente identificada como S. speluncae.

Palavras chave: Scytalopus speluncae, espécie nova, Brasil, taxonomia

Introduction

The genus Scytalopus Gould, 1836, is one of the most interesting and taxonomicallycomplex of all suboscine passerine birds. Its members are small, usually gray, associatedwith mountains, and range from Costa Rica to southernmost Argentina (Whitney 1994;Krabbe & Schulenberg 1997). Populations of Scytalopus are highly susceptible to localisolation and many species have proved difficult to detect because of their tinydistributions. The vast majority of species are morphologically very similar, and in manyinstances may be distinguished only by their vocalizations and molecular data. Only veryrecently have ornithologists gained sufficient experience and confidence to conduct much-needed revisions of species limits within the genus. Consequently, the number of speciesrecognised within Scytalopus has risen steadily, from 11 in 1970 to about 40 nowadays,with two additional species, both from Colombia, described in 2005 alone (Cuervo et al.2005; Krabbe et al. 2005).

The species-group Scytalopus speluncae occurs in south-eastern Brazil and extremenortheastern Argentina and has been subject to reasonable scrutiny over the last 50 years(e.g. Sick 1960; Vielliard 1990; Krabbe & Schulenberg 2003), resulting in the descriptionof three new species, Scytalopus novacapitalis Sick, 1958, Scytalopus iraiensisBornschein et al., 1998, and Scytalopus pachecoi Maurício, 2005.

Scytalopus speluncae was based on a specimen described as having been taken at SãoJoão del Rei, Minas Gerais, Brazil and having the throat and the central breast evidentlywhitish (“devident blanchâtre vers le milieu de la gorge et de la poitrine”, Ménétriés 1835:527). Since then, however, this name has rather surprisingly been attributed to theuniformly slate gray Mouse-colored Tapaculo of the Serra do Mar and vicinity in theeastern coastal Atlantic Forest of Brazil (Fig. 1, see also Krabbe & Schulenberg 2003). The first author to identify the eastern Atlantic Forest birds as S. speluncae was Ihering(1907), who applied this name as a replacement for Scytalopus sylvestris Taczanowski,1874, used by Miranda-Ribeiro (1905). Later, Miranda-Ribeiro (1923) would alsoassociate the names Scytalopus magellanicus (Gmelin, 1789) and Scytalopus niger(Swainson, 1838) with specimens from Itatiaia and Nova Friburgo, in the state of Rio deJaneiro. All these names (except S. speluncae) were considered inapplicable by Chapman(1915), Cory and Hellmayr (1924), and Holt (1928) as their types are all from the Andeanregion.

Application of the name S. speluncae in all subsequent literature to the Serra do Mar

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1271ZOOTAXApopulations may in part be attributed to the fact that, since the 1830s, no species of

Scytalopus had been recorded in the environs of São João del Rei (Vielliard 1990). For thisreason, some authors questioned the validity of São João del Rei as the correct typelocality (Sick 1960; Maurício 2005), a judgment reinforced by obvious problemsassociated with the type locality of other specimens collected by Ménétriés during hiscomparatively brief sojourn in Brazil (Pacheco 2004). Vielliard (1990) suggested thatenvironmental degradation might explain what he mistakenly thought to be the modern-day absence of the species in this region, but he was also forced to admit that São João delRei appears somewhat inland of the currently accepted distribution of the slate gray form.

During field work in the Chapada Diamantina, Bahia, in the northernmost part of theEspinhaço geological complex, we collected some specimens of Scytalopus in December1995 and October 1996 that are somewhat intermediate between the whitish-gray S.novacapitalis and the dark gray coastal specimens traditionally identified as S. speluncae.The Chapada Diamantina lies within the same geological complex as São João del Rei,although the localities are situated at opposite extremes of this range. At that time, it wasevident that analysis of the holotype and of fresh material from the type locality of S.speluncae would be needed to resolve the identification of the new specimens, so theiridentity was left unresolved and they were listed simply as "Scytalopus sp." in Parrini et al.(1999: 94).

As part of his study, Maurício (2005:18) analysed our specimens from the ChapadaDiamantina, also referring to them as Scytalopus sp., which treatment he also applied toother specimens from the Espinhaço range in Minas Gerais (see also Krabbe &Schulenberg 2003: 771). He considered that several populations of Scytalopus in easternBrazil almost certainly await description, presumably as species. Of paramountimportance, however, is the resolution of the correct application of the name Scytalopusspeluncae, the senior species name of the superspecies, as none of the recent taxonomicrevisions of east Brazilian Scytalopus (Ridgely & Tudor 1994; Bornschein et al. 1998;Krabbe & Schulenberg 2003; Maurício 2005) has included a reanalysis of the holotype,which is housed at the Zoological Institute (Russian Academy of Sciences), St. Petersburg.

Our study of this case corroborates the conclusions of Krabbe and Schulenberg (2003)and Maurício (2005), who accepted the existence of two species within the Scytalopusspeluncae species-group in southeastern Brazil. According to these authors and our ownanalysis, the populations of Scytalopus from the Espinhaço range of Minas Gerais andBahia differ from those of the Serra do Mar and vicinity. These two species can bedistinguished by the following characters. The Serra do Mar species is uniformly BlackishNeutral Gray (82–83, of Smithe 1975) whereas the Espinhaço species is Medium to LightNeutral Gray (86) with some whitish on the abdominal area, as well as having the rump,thighs and vent black with rufous fringes to the feathers (Fig. 1). Their vocalizations arealso quite different.

However, our analysis of the holotype, the two new topotypical specimens, and the

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1271ZOOTAXA original description of Scytalopus speluncae strongly suggests that this name must be

applied to the pale gray form with rufous-barred thighs that is found in Minas Gerais. Thispaper redescribes Scytalopus speluncae, correcting its application, and describing, as anew species, populations from the Serra do Mar previously ‘misidentified’ as S. speluncae.

Methods

We analyzed two recently collected topotypes of S. speluncae collected by MAR and RS,

in September 2005 at São João del Rei, Minas Gerais (21o04’16.8’’S, 44o20’19.4’’W), andthe holotype (ZISP 145251) and type description (including original plate) of Scytalopusspeluncae. For diagnosis of taxa we adopted standard English terms, with occasionalreference to some of the Latin terms of Baumel et al. (1993). We also examined specimensof all species of the complex held at the Museu Nacional, Universidade Federal do Rio deJaneiro (MNRJ), Museo Argentino de Ciencias Naturales, Buenos Aires (MACN), Museude Zoologia da Universidade de São Paulo (MZUSP), and American Museum of NaturalHistory, New York (AMNH), including the holotype and paratypes of Scytalopusnovacapitalis (MNRJ) and S. iraiensis (MNRJ). The specimens of S. pachecoi are fromMisiones, Argentina (MACN and AMNH). Specimen analysis focused on plumagecoloration (using Smithe 1975 as a basis for colors) and morphometric characters (lengthof wing, tail, and exposed culmen). Morphometric analysis included only descriptivestatistics of the species as a whole.

Sound-recordings of the Scytalopus speluncae species-group were obtained fromprivate collections (Vitor Torga and Ricardo Parrini), and were used only to complementdescriptions of the taxa. The vocalizations (n=3) from São João del Rei (Vitor Torga) wererecorded in the same place that we collected the new specimens, and were used to attractthem. For the acoustic analysis, the programs Avisoft and SoundRuler (version 0.9.4.)were used. Recordings of all vocalizations are stored at the Museu Nacional (see Appendix2).

Results

Restriction of Scytalopus speluncae Ménétriés, 1835

Type locality. Despite Vielliard's (1990) assertion of absence, this species is apparentlyvery common at the type locality of São João del Rei. Two topotypes were collected in twodegraded woodlots that agree quite well with Ménétriés’s description of the area in whichthe type was collected (“courant à terre et voltigeant sur les petits buissons, à l’entréed’une grotte calcaire près de St. João del Rey, Minas Gerais”).

Мénétriés’s diary notes are also in perfect accordance with those of the leaderLangsdorff (Mikulinskii 1995), who places the expedition at Barbacena, Minas Gerais,

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1271ZOOTAXAfrom 1–4 June, when, on 5 June, it was decided to proceed to São João do Rei. On 6 June

he remarked that “We have passed by Fazenda Barroso near the brook Ribeyran , ..reached the River Ribeyran-du-kayeyra [?] which falls into the River Mortis [Rio dasMortes]…., and then reached the fazenda of Lorenso Pintu” (p.33); on 7 June Langsdorffleft “Invernadu” for São João del Rei, noting that "In Barbaсena, we have receivedinformation about a very remarkable grotto situated in the environs of São João del Rey orpossibly beside the River Elvas. . . . I supposed that, as we agreed, I shall meet there mytraveling companions at 12; …. I galloped forward through the Invernadu [valley]…. At10.30 I stopped ... and heard that my companions [including Мénétriés] spent the nightthere and went to the grotto" (p.34). "I arrived there at 12 and found there Ridel, Мénétriésand Rugendas. We left the grotto at about 3 p.m. and arrived at the town of São João delRey at about 5 p.m.…." (p. 35). It is obvious from these remarks, which we have translatedfrom the Russian, that Мéntriés was collecting very close to São João del Rei and waswithin two hours ride of this town. The new topotypes were collected in gallery forest ofthe Rio das Mortes, which river is also referred to by Langsdorff, and is very close to SãoJoão del Rei.

Maurício (2005) dismissed São João del Rei as the type locality of Scytalopusspeluncae because of historical “problems surrounding the origins of Ménétriés’ types”,reflecting the conclusions of Pacheco (2004), who showed several of Ménétriés’ typelocalities to be improbable.

In the present instance, however, there would appear to be no grounds for alteringМénétriés clear designation, which has been substantiated by the modern collection ofspecimens agreeing with the holotype near the type-locality.

The holotype of Scytalopus speluncae. Мénétriés (1835: 527) made no mention of therufous and black stripes on the tips of the rump and flank feathers in the originaldescription of S. speluncae, nor were they illustrated (pl. 13, fig. 1). It seems likely thatthese parts were already damaged either in the collection or preparation so that Мénétriéswould not have been aware of such features, in 1835, some ten years after the specimenwas collected. That vestiges of these characters still remain on the holotype makes theiromission from the plate and type description less important.

The holotype of S. speluncae is an adult male, as indicated by the plain upper wingcoverts, which lack black markings (Figs. 2–3). Unfortunately, the belly feathers areseverely damaged, and almost all of the vent and thigh feathers are in such poor conditionthat it is almost impossible to discern their original color, except that of the blackish base(ground) color to these tracts (Fig. 4). However, this is not the case with the rump feathers,which are buff with dark bars (Fig. 5). A careful analysis of the holotype by VL furtherrevealed some tiny remnants of buff color on one feather on the left thigh, and on threefeathers on the right flank.

The type description is also very important in establishing the original color of theholotype. The author states that the throat and the middle of the breast of his Scytalopus

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1271ZOOTAXA speluncae are whitish (see Introduction), a description that could never be applied to the

Blackish Neutral Gray (82–83) Scytalopus from the Serra do Mar. The plate thataccompanied the type description (Fig. 6) also shows considerable whitish on the ventralsurface, despite not showing the buff feathers found on the flanks and rump and, evidentlymore sparsely, on the thighs of the holotype. In sum, the type description points to a gray-colored bird, with a partially whitish venter, and close examination of the holotype showsthat it also has rufous-barred rump feathers, and at least some buff on the thighs and vent.The topotypes are also pale gray with a whitish belly and buff-barred vent, thighs, andrump (Figs. 7–9).

FIGURE 1. Comparison between two specimens of Scytalopus. Left: the slate gray Mouse-coloredTapaculo, specimen from Nova Friburgo, in the Serra do Mar range. Right: a specimen belonging tothe lighter form from Chapada Diamantina, in the Espinhaço Range.

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FIGURES 2–5. Holotype of Scytalopus speluncae (ZISP 145251): 2, dorsal view; 3, lateral view;4, ventral view; 5, uropygium showing buff-barred black feathers.

Maurício (2005) stated, based on the original plate, that Ménétriés (1835) must havedescribed an “unbarred, adult bird”. He also considered that Chrostowski (1921), who was

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1271ZOOTAXA one of the few persons prior to ourselves to have directly analysed the holotype, had erred

in describing the presence of rufous fringes to the rump feathers, as this supposedly couldnot be corroborated by either the original description or by the photographs of the holotypethen available (which are of very poor quality; pers. obs. all authors). Now, however, wecan state with assurance that Chrostowski was correct in his assertion regarding theholotype’s features.

FIGURE 6. The original plate of Scytalopus speluncae (Ménétriés 1835, pl. 13, fig. 1).

It is evident that current application of the name Scytalopus speluncae to the dark grayspecimens from the Serra do Mar is erroneous and the name must be used instead for thepopulations of the Espinhaço range. Surprisingly, we have been unable to locate any otherapplicable name for the well-known Serra do Mar species. Therefore we propose a nameto this species here and redescribe S. speluncae, which name hereafter we use solely forthe Espinhaço population.

Characterization of species

Scytalopus notorius new species

Diagnosis: Male easily diagnosed from all other Brazilian Scytalopus by its homogeneousBlackish Neutral Gray (82) coloration, being slightly darker on the upperparts. It is furtherdistinguishable from the paler species, Scytalopus speluncae, S. pachecoi, and S.novacapitalis by the gray feathers of the flanks, thighs, vent and upper tail coverts, whichare quite unlike the buff-fringed blackish feathers of those three species. S. notorius differsfrom S. iraiensis by its gray versus blackish upperparts and in lacking any contrastbetween the dorsal and ventral areas. Males and females of Scytalopus iraiensis furtherdiffer from S. notorius in having blackish flanks, thighs, and vent, and broader tailfeathers, as well as in behavioral differences (Figs. 7–10).

Description of the holotype: Adult male (MNRJ 36652) (Figs. 7–9), collected at

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1271ZOOTAXANova Friburgo, Rio de Janeiro, in September 1989. Bill black; forehead, crown, occiput,

nape, mantle, rump, upper tail coverts, wing coverts and dorsal surface of rectricesBlackish Neutral Gray (82); lores, auriculars, chin, throat, upper breast, abdomen, flanks,crural feathers, crissum, primary and secondary under wing coverts, under tail coverts, andventral surface of rectrices Dark (83) to Blackish Neutral Gray (82). Tarsus and digits palebrown (label data). Exposed culmen, 11.7 mm; tail, 41.0 mm; wing (chord), 50.2 mm;body mass, 16.0 g.

Description of the paratypes (Fig. 10): Adult female (MNRJ 36653), collected atNova Friburgo, Rio de Janeiro, in September 1989. Maxilla black and mandible brown(label data); forehead Blackish Neutral Gray (82); crown, occiput, nape and mantleBlackish Neutral Gray, shaded Brownish Olive (29); rump, upper tail coverts and dorsalsurface of rectrices Brownish Olive, fringed black; upper secondary coverts BrownishOlive with transverse subterminal black patch and an Army Brown (219B) apical area;primaries and secondaries Homogeneous Sepia (119); lores, auriculars, throat and upperbreast Dark Neutral Gray (83); feathers of the flanks, crural feathers, crissum and undertail coverts brown with transverse black patches; tarsus and digits yellowish brown (labeldata). Exposed culmen, 11.7 mm; tail, 40.5 mm; wing (chord), 51.1 mm; body mass, 15.8g.

Subadult female (MNRJ 36655): collected at Nova Friburgo, Rio de Janeiro, inSeptember 1989. Maxilla black and mandible brown (label data); forehead BlackishNeutral Gray; crown, occiput, nape, mantle and dorsal surface of rectrices BlackishNeutral Gray shaded Brownish Olive; upper tail coverts and rump brown with twotransverse black patches; lesser secondary coverts Brownish Olive with a transversesubterminal black patch and an Army Brown apical area; primaries and secondaries Sepiawith different degrees of black at their tips and, sometimes, tiny white notches; lores,auriculars, throat and upper breast Dark Neutral Gray; flanks, crural feathers, crissum andunder tail coverts brown with transverse black patches; tarsus and digits yellowish brown(label data). Exposed culmen, 11.9 mm; tail, 40.0 mm; wing (chord), 52.9 mm; body mass,15.0 g.

Young female (MNRJ 36656): collected at Nova Friburgo, Rio de Janeiro, inSeptember 1989. Maxilla black and mandible brown (label data); forehead BlackishNeutral Gray shaded Brownish Olive; crown, occiput, nape, mantle and dorsal surface ofrectrices Brownish Olive; rump and upper tail coverts brown with two transverse blackpatches; lesser secondary coverts Brownish Olive, similar to adult female, with atransverse subterminal black patch and an Army Brown apical area; primaries andsecondaries Sepia; lores, auriculars, throat and upper breast Medium Neutral Gray (84);abdomen, flanks, crural feathers, crissum and under tail coverts Buff (124) with transverseblack patches; tarsus and digits brown (label data). Exposed culmen, 12.2 mm; tail, 41.0mm; wing (chord), 48.8 mm; body mass, 14.5 g.

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FIGURES 7–9. From left to right in each figure (all males): paratype of Scytalopus iraiensis;holotype of S. notorius, new species; topotype of S. speluncae; and holotype of S. novacapitalis; 7,ventral view; 8, lateral view; 9, dorsal view.

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FIGURE 10. Lateral view of females: holotype of Scytalopus iraiensis; topotype of S. speluncae;paratype of S. notorius.

Morphometry : See Table 1.Vocalizations: The song phrases are of very variable duration and consist of regular

repetition of the same notes (Fig. 11), whereas the mean duration of the intervals betweenthe different notes is 0.15 seconds. Thus a bird sings approximately five notes per second.Each note lasts for a mean of 0.05 seconds, and first ascends before descending (Fig. 12),although the overall frequency varies only between 2.6 and 2.8 kHz, and the dominantfrequency between just 5.3 and 5.4 kHz. The alarm-call consists of a single monosyllabicnote with a mean of 0.1 seconds duration, at 1.8 kHz, with the dominant frequency 3.7 kHz(Fig. 13).

Type locality: Nova Friburgo, Rio de Janeiro, Brazil. The holotype and paratypes

were collected at Três Picos (22o20'10.02"S, 42o42'44.31"W), a locality now includedwithin the Parque Estadual dos Três Picos. The specimen labels do not record the altitudewhere the specimens where taken, but the collector (G. Luigi, in litt.) has informed us thatthey were taken between 1200 and 1700 m.

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FIGURES 11–13. Spectrograms of the vocalizations of Scytalopus notorius, new species(Teresópolis at Serra do Mar, Ricardo Parrini). 11, complete song; 12, ten elements of the samesong of Scytalopus speluncae in a different scale; 13, call.

Geographic range: Scytalopus notorius ranges from central-west Espírito Santo, onthe border with Minas Gerais (in the Serra do Caparaó) through Rio de Janeiro south toRio Grande do Sul. Maurício (2005) considered that this form (treated by him as S.speluncae) might be best regarded as consisting of two different species, one occurringfrom São Paulo northwards, and the other from São Paulo southwards, a conclusion thatappears highly plausible, but for which much additional evidence needs to be garnered.

Etymology: From Medieval Latin nōtōrius, well-known, from Latin nōtus, known,

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1271ZOOTAXAreflecting the fact that the new species represents a long-known population that has been

inappropriately identified as S. speluncae for over 100 years.

FIGURES 14–16. Spectrograms of the vocalizations of Scytalopus speluncae (São João del Rei,Vitor Torga). 14, complete song; 15, five elements of the same song of Scytalopus speluncae in adifferent scale; 16, call.

Scytalopus speluncae Ménétriés, 1835

Diagnosis: Adult male easily diagnosed from Scytalopus notorius by its paler ventral graycolor (84 to 85, rather than 83), Pale Neutral Gray (86) to whitish abdomen (belly), rather

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1271ZOOTAXA than Dark Neutral Gray (83) of S. notorius, and by the buff-fringed black feathers of the

flanks, thighs, vent and upper tail coverts. It differs from S. iraiensis by its Dark NeutralGray color (83), unlike the blackish upperparts of the latter species, which is only knownfrom wetland habitats, and again by the buff-fringed black feathers of the flanks, thighs,vent and upper tail coverts. Morphological diagnosis in relation to S. novacapitalis and S.pachecoi is very difficult. The two topotypes of S. speluncae are paler than mostspecimens of S. pachecoi and slightly darker then the holotype and paratypes of S.novacapitalis. All three species have buff-fringed black feathers in flanks, thighs, vent andupper tail coverts, but S. novacapitalis has more whitish on the abdomen, and S. pachecoi,in general, has a pale gray abdomen.

Description of the holotype (Figs. 2–5): Adult specimen (ZISP 145251) withauriculars, throat and upper breast close to Medium Neutral Gray (84) but with a palebrown tint. Upperparts close to Dark Grayish Brown (20) or Vandyke Brown (121). Itshould be mentioned that the holotype is almost 180 years old, and thus the original colorof the upper and under parts may have acquired a somewhat brownish tone (foxing). Mostof the feathers of the abdomen, flanks, crural feathers and under tail coverts are damagedand have lost their structure and color, but there are still small remnants of Clay Color(123B) on one feather of the left thigh, and on three feathers on the right flank (ilia); uppertail coverts Clay Color fringed black.

Description of the topotypes: Adult male (MNRJ 44019, Figs. 7–10); collected at

São João del Rei, Minas Gerais, at the right bank of Rio das Mortes (21o04’16.8’’S,

44o20’19.4’’W) in September 2005. Mandible blackish with gray tomia; crown, occiput,nape, greater primary coverts, and mantle Dark Neutral Gray (83); rump, upper secondarycoverts, and dorsal surface of rectrices Dark Brownish Olive (129); upper tail coverts Buffto Clay Color (124–123B) with transverse black marks; lores Light Neutral Gray (85) withsmall black spots at the tips of the feathers; auriculars, throat, upper breast, secondary andunder primary coverts Light Neutral Gray (85); wrist coverts (tectrices propatagiventrales) and abdomen whitish, bordered distally with Pale Neutral Gray (86); abdomen,flanks, crural feathers and under tail coverts Clay Color with transverse black marks,tarsus and digits Olive-Yellow (52). Exposed culmen, 10.0 mm; tail, 40.0 mm; wing(chord), 48.9 mm; mass, 16.5g.

Adult female (MNRJ 44018, Fig. 2); collected at São João del Rei, Minas Gerais, inSeptember 2005. Mandible blackish with gray tomia; forehead Dark Neutral Gray;occiput, nape and mantle; upper secondary coverts and greater primary coverts DarkBrownish Olive with varying degrees of black at their distal end; dorsal surface ofrectrices Dark Brownish Olive; rump and upper tail coverts Clay Color with transverseblack marks; lores Light Neutral Gray with small black marks at the tips of the feathers;auriculars, throat, upper breast light neutral gray; secondary under coverts, primary undercoverts and wrist coverts (tectrices propatagi ventrales) Buff (124) with narrow blackmarks; abdomen whitish, distally bordered Pale Neutral Gray; flanks, crural feathers,

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1271ZOOTAXAcrissum and under tail coverts Clay Color with transverse black marks; tarsus and digits

Olive-Yellow (52). Exposed culmen, 10.7 mm; tail, 40.0 mm; wing (chord), 44.3 mm;mass, 13.5 g.

Geographic variation: The four specimens analyzed by us from the northern end ofthe Espinhaço Range, in the Chapada Diamantina (Fig. 1), are somewhat different fromthe topotypes. These specimens lack the whitish ventral area and could be confused with S.pachecoi (Maurício 2005) from Misiones (Argentina), or Rio Grande do Sul and SantaCatarina (Brazil). The upperparts of these specimens are Dark Neutral Gray (83) and theunderparts Medium Neutral Gray (84), with buff-marked black feathers on the thighs,flanks, and crissum.

FIGURE 17. Map showing the distribution of the Scytalopus speluncae species-group (based onthe specimens examined in this study).

Morphometry : See Table 1.Vocalizations: The song phrases are variable in their duration and, like those of S.

notorius, consist of regular repetitions of the same notes (Fig. 14). However, the mean

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1271ZOOTAXA time between notes is 0.38 seconds. Thus the bird typically sings just 2.5 notes per

second. Each note is very brief, lasting on average just 0.03 seconds, and descending inoverall frequency (Fig. 15), whereas the fundamental frequency varies between 1.8 and2.0 kHz, and the dominant frequency between 3.5 and 4.1 kHz. Like S. notorius, thealarm-call consists of a single monosyllabic note which lasts a mean of 0.07 seconds, witha fundamental frequency of 3.5 kHz and a dominant frequency of 5.7 kHz (Fig. 16). Thesong of S. speluncae is easily distinguished from that of S. notorius by its pace, which isapproximately half that of S. notorius (2.5 notes per second vs. 5.0 notes per second),whilst the modulation describes an ascent, then descent in S. notorius but solely descendsin S. speluncae. On the other hand, the song of S. speluncae is indistinguishable from thatof S. pachecoi by pace, as S. pachecoi delivers 2.7 notes per second (Maurício 2005:13)and S. speluncae 2.5 notes per second, and both descend in frequency. The call of S.speluncae is also identical to the call described by Maurício (2005) for S. pachecoi. Inrelation to S. novacapitalis (based on Vielliard 1990 and Maurício 2005) both S. notoriusand S. speluncae possess distinctive characters, as the song of S. novacapitalis consists onaverage of just 1.1 notes per second, which is considerably fewer than either S. notorius orS. speluncae, whereas the duration of each note lasts on average 0.1 seconds, as opposed toeither S. speluncae or S. notorius, which never exceeded 0.05 seconds in any sampleanalyzed.

Type locality: São João del Rei, Minas Gerais, is the type locality by originaldesignation.

TABLE 1. Mensural data for the Scytalopus speluncae species group. Mean, standard deviation(SD), minimum value (min), maximum value (max), number of specimens examined (n), male (m)and female (f).

Geographic range: Known thus far only from the opposite ends of the Espinhaço

Bill length (mm) Wing length (mm) Tail length (mm)

mean±SD min–max n mean±SD min–max n mean±SD min–max n

S. notorius m 11.29±0.52 10.50–12.10 10 49.32±2.58 44.00–54.00 10 44.00±2.12 41.00–47.50 9

f 11.56±0.58 10.85–12.20 5 49.53±2.42 47.15–52.90 5 41.51±1.57- -40.05–44.00 5

S. speluncae m 12.00±0.95 10.95–12.80 3 51.50±3.70 48.05–55.40 3 44.00±4.58 40.00–49.00 3

f 11.33±0.81 10.75–11.90 2 48.33±3.56 44.35–51.20 3 40.00±2.00 38.00–42.00 3

S. novacapitalism - - 0 50.65 50.65 1 42.50 42.50 1

f - - 0 - - 0 - - 0

S. iraiensis m 10.80 10.80 1 50.20 50.20 1 39.50 39.50 1

f 10.90 10.90 1 46.15 46.15 1 39.00 39.00 1

S. pachecoi m 11.65±0.99 10.95–12.35 2 50.03±0.25 49.85–50.20 2 40.75±1.77 39.50–42.00 2

f 11.25±0.74 10.40–11.75 3 44.33±0.34 43.95–44.60 3 37.67±2.08 36.00–40.00 3

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1271ZOOTAXArange at São João del Rei, Minas Gerais, in the south, and Chapada Diamantina, Bahia, in

the north (Fig. 17).

Acknowledgments

We are grateful to the Fundação Carlos Chagas Filho de Amparo à Pesquisa do Estado doRio de Janeiro (FAPERJ) for their support of M. A. Raposo (proc. Instalação E-26/170.871/2003 and Primeiros Projetos E-26/ 170.642/2004) and Renata Stopiglia (BolsaNota 10, E-26/ 150.404/2005). Ulisses Caramaschi and Dante Martins Teixeira were veryhelpful with nomenclatural issues. Vitor Torga was the first to rediscover S. speluncae inSão João del Rei and, along with Ricardo Parrini, supplied vocal material of relevant taxa.Claydson P. de Assis and Liliane Seixas assisted our fieldwork. The Instituto Brasileiro deMeio Ambiente e dos Recursos Naturais Renováveis (IBAMA) has provided collectingpermits to Marcos A. Raposo. The manuscript benefited very much from comments ofStorrs Olson (Associate Editor), Niels Krabbe, Herculano Alvarenga and two anonymousreferees.

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1271ZOOTAXAAppendix 1. Specimens examined.

Scytalopus notorius new species: BRAZIL: Rio de Janeiro : Casa Queimada, Serra do Caparaó,M, MNRJ 14202. Serra do Caparaó, M, MNRJ 26267, 26281, 27035, 27036. Fazenda Toledo,Três Picos, Nova Friburgo, M, MNRJ 36652 (holotype); F, MNRJ 36653, 36655, 36656(paratypes). Campos de Itatiaia, M, MZUSP 6121. Parque Nacional de Itatiaia, M, MZUSP34381, 34804, 34806, 36347; F, MZUSP 34805, 34807. São Paulo: Alto da Serra, MZUSP4836.

Scytalopus speluncae: BRAZIL: Minas Gerais : São João del Rei, M, ZISP 145251 (holotype);MNRJ 44019; F, MNRJ 44018. Bahia: Ibicoara, Chapada Diamantina, M, MNRJ 42741; F,MNRJ 42742. Bonito, Chapada Diamantina, M, MNRJ 43053; F, MNRJ 43054.

Scytalopus novacapitalis: BRAZIL: Goiás: Distrito Federal, MNRJ 27905 (paratype); M, MNRJ27906 (holotype).

Scytalopus iraiensis: BRAZIL: Paraná : Quatro Barras, F, MNRJ 43378 (holotype); M, MNRJ43379 (paratype).

Scytalopus pachecoi: ARGENTINA : Misiones, Arroyo Uruguai, M, AMNH 795288, 795289;MACN 33037, 36868, 36870. F, AMNH 771239, 771240, 771242; MACN 33038, 33039,36869, 33623.

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1271ZOOTAXA Appendix 2. Recordings examined

Scytalopus notorius new species: Brazil: Rio de Janeiro, Itatiaia (2 songs and 2 calls); ParqueEstadual do Desengano (1 song); Serra do Caparaó (1 song); Teresópolis (1 song and 1 call).

Scytalopus speluncae: Brazil: Minas Gerais, São João do Rei (3 songs and 2 calls); Bahia,Lençóis (2 songs and 1 call); Ibicoara (3 songs and 2 calls).